STABILIZING SELECTION DETECTED FOR BRISTLE NUMBER IN DROSOPHILA MELANOGASTER

Stabilizing selection, which favors intermediate phenotypes, is frequently invoked as the selective force maintaining a population's status quo. Two main alternative reasons for stabilizing selection on a quantitative trait are possible: (1) intermediate trait values can be favored through the causal effect of the trait on fitness (direct stabilizing selection); or (2) through a pleiotropic, deleterious side effect on fitness of mutants affecting the trait (apparent stabilizing selection). Up to now, these alternatives have never been experimentally disentangled. Here we measure fitness as a function of the number of abdominal bristles within four Drosophila melanogaster lines, one with high, one with low, and two with intermediate average bristle number. The four were inbred nonsegregating lines, so that apparent selection due to pleiotropy is not possible. Individual fitness significantly increased (decreased) with bristles number in the low (high) line. No significant fitness-trait association was detected within each intermediate line. These results reveal substantial direct stabilizing selection on the trait.     

EQUILIBRIUM ANALYSIS OF SEXUAL SELECTION IN DROSOPHILA MELANOGASTER

Several models for sexual selection, both by male-male competition and female choice, predict that a character which covaries with mating success should be near an equilibrium where the intensity of sexual selection opposes viability selection. This prediction was used to design experiments for estimating the intensity of sexual and viability selection on wing length in a recently captured population of Drosophila melanogaster. Observations of matings by males color-marked for wing length indicated that the standardized sexual selection differential on wing length was 0.24 under a wide range of effective sex ratios. After estimating the heritability of wing length to be 0.62, the expected standardized response due to sexual selection was calculated as 0.15 (SE = 0.15). The response due to viability selection was then estimated by comparing wing lengths of progeny of flies that had been randomly mated, thereby preventing sexual selection, with progeny of flies that had been allowed to acquire mates in a mass-mating chamber. The results support an equilibrium model in that the standardized response due to viability selection (?0.31, SE = 0.08) was opposite in sign and similar in magnitude to the estimated response due to sexual selection. Observations of females orienting in front of males which differed in wing length indicated that the mating advantage accruing to long-winged males was not due to female choice. Instead, male-male competition in which the larger of two randomly chosen males succeeded in mating, explains the observed sexual selection. An experimental analysis of genotype-environment interaction revealed that larval density had a nonlinear effect on mean wing length within sibships. If a population is displaced from equilibrium, therefore, the evolutionary trajectory of mean wing length will depend both on the intensity of selection and the environment in which that selection is operating.     

CROSS-RESISTANCE FOLLOWING ARTIFICIAL SELECTION FOR INCREASED DEFENSE AGAINST PARASITOIDS IN DROSOPHILA MELANOGASTER

An increase in resistance to one natural enemy may result in no correlated change, a positive correlated change, or a negative correlated change in the ability of the host or prey to resist other natural enemies. The type of specificity is important in understanding the evolutionary response to natural enemies and was studied here in a Drosophila-paxasitoid system. Drosophila melanogaster lines selected for increased larval resistance to the endoparasitoid wasps Asobara tabida or Leptopilina boulardi were exposed to attack by A. tabida, L. boulardi and Leptopilina heterotoma at 15°C, 20°C, and 25°C. In general, encapsulation ability increased with temperature, with the exception of the lines selected against L. boulardi, which showed the opposite trend. Lines selected against L. boulardi showed large increases in resistance against all three parasitoid species, and showed similar levels of defense against A. tabida to the lines selected against that parasitoid. In contrast, lines selected against A. tabida showed a large increase in resistance to A. tabida and generally to L. heterotoma, but displayed only a small change in their ability to survive attack by L. boulardi. Such asymmetries in correlated responses to selection for increased resistance to natural enemies may influence host-parasitoid community structure.     

DISRUPTIVE SELECTION ON BODY WEIGHT IN DROSOPHILA MELANOGASTER

Disruptive selection can have two consequences in a population: the rise of polymorphism or the divergence of the population. Divergence creates subpopulations which can be sexually isolated. Previous studies have shown that these consequences depend on the pattern of mate choices by the individuals. We have studied the effects of different patterns of mating (free mating, enforced random mating and enforced assortative mating) in a population of Drosophila melanogaster subjected to disruptive selection on body weight. Disruptive selection increased the weight dispersion in all mating patterns but brought about divergence within the population only when it occurred in association with enforced assortative mating (without complete sexual isolation between the subpopulations). When mating was free, the dispersion, measured by change in the coefficient of variation, was only somewhat larger than with enforced random mating: thus the principal consequence of disruptive selection was the rise of polymorphism in the population.     

DIRECT SELECTION ON LIFE SPAN IN DROSOPHILA MELANOGASTER

An important issue in the study of the evolution of aging in Drosophila melanogaster is whether decreased early fecundity is inextricably coupled with increased life span in selection experiments on age at reproduction. Here, this problem has been tackled using an experimental design in which selection is applied directly to longevity. Selection appeared successful for short and long life, in females as well as males. Progeny production of females selected for long life was lower than for short-lived females throughout their whole life. No increase of late-life reproduction in long-lived females occurred, as has been found in selection experiments on age at reproduction. This discrepancy is explained in terms of the inadequacy of the latter design to separate selection on life span from selection on late-life fecundity. Moreover, starvation resistance and fat content were lower for adults selected for short life. In general, the data support the negative-pleiotropy–disposable-soma theory of aging, and it is hypothesized that the pleiotropic allocation of resources to maintenance versus to reproduction as implicated in the theory might involve lipid metabolism. It is argued that further research on this suggestion is urgent and should certainly comprise observations on male reproduction because these are for the greater part still lacking. In conclusion, the longevity of D. melanogaster can be genetically altered in a direct-selection design, and such an increase is accompanied by a decreased general reproduction and thus early reproduction.     

DIRECTIONAL AND STABILIZING DENSITY-DEPENDENT NATURAL SELECTION FOR PUPATION HEIGHT IN DROSOPHILA MELANOGASTER

Six populations of Drosophila melanogaster have been kept at extreme population densities, three high and three low, for 175 generations. Larvae from the high density populations pupate 50%-100% higher than larvae from the low density populations. At high larval test densities there is both a directional and a stabilizing component to selection, with viabilities ranging from 0.14 to 0.992, depending on the choice of pupation site. The directional component is stronger on the populations which have evolved at low densities, while the stabilizing component is stronger on the populations which have evolved at high densities. There is no indication that the evolution of this trait, in response to density, has altered its phenotypic plasticity.     

ALLOZYMIC DIFFERENTIATION IN RESPONSE TO LABORATORY DEMOGRAPHIC SELECTION OF DROSOPHILA MELANOGASTER

Drosophila melanogaster populations that exhibit constrasting life histories as a result of laboratory selection were compared at several potentially relevant enzyme loci. Selection regimes included postponed reproduction, accelerated development, and intermediate generation time. Each selection regime was represented by fivefold replicated populations maintained for between 50 and 500 generations. For each population, allele frequencies were calculated from frequencies of electrophoretically distinguishable allozymes of alcohol dehydrogenase, α-glycerol-3-phosphate dehydrogenase, phosphoglucomutase, and CuZn-superoxide dismutase. Based on allozyme frequency changes consistent across replicate populations, two of the studied loci responded to both selection for postponed reproduction and selection for accelerated development. The responses to contrasting selection regimes were in opposing directions, suggesting antagonistic pleiotropy.     

RESISTANCE OF GENETIC CORRELATION STRUCTURE TO DIRECTIONAL SELECTION IN DROSOPHILA MELANOGASTER

The genetic covariance and correlation matrices for five morphological traits were estimated from four populations of fruit flies, Drosophila melanogaster, to measure the extent of change in genetic covariances as a result of directional selection. Two of the populations were derived from lines that had undergone selection for large or small thorax length over the preceding 23 generations. A third population was constituted using flies from control lines that were maintained with equivalent population sizes as the selected lines. The fourth population contained flies from the original cage population from which the selected and control lines had been started. Tests of the homogeneity of covariance matrices using maximum likelihood techniques revealed significant changes in covariance structure among the selected lines. Prediction of base population trait means from selected line means under the assumption of constant genetic covariances indicated that genetic covariances for the small population differed more from the base population than did the covariances for the large population. The predicted small population means diverged farther from the expected means because the additive genetic variance associated with several traits increased in value and most of the genetic covariances associated with one trait changed in sign. These results illustrate that genetic covariances may remain nearly constant in some situations while changing markedly in others. Possible developmental reasons for the genetic changes are discussed.     

CORRELATED RESPONSES IN LIFE-HISTORY TRAITS TO ARTIFICIAL SELECTION FOR BODY WEIGHT IN DROSOPHILA MELANOGASTER

Drosophila melanogaster that had been successfully selected on rich and poor larval medium for increased and decreased fresh weight at eclosion were tested on an intermediate medium for correlated responses in longevity, fertility, and hatchability. Larger flies laid more eggs early in life and lived shorter lives than smaller flies, which not only lived longer but also laid more eggs later in life. This supports the notion of a mortality cost of reproduction in Drosophila. The total number of eggs laid per lifetime did not differ between the two groups. The percentage of offspring hatched started at normal levels (about 50% of eggs laid), then declined rapidly in large flies. In small flies, hatchability started at a lower level early in life (40-65%), but declined less rapidly, and later in life was higher than the hatchability of eggs laid by larger flies.     

SEXUAL SELECTION ACCELERATES THE ELIMINATION OF A DELETERIOUS MUTANT IN DROSOPHILA MELANOGASTER

Although theory indicates that indirect genetic benefits through mate choice should be widespread, empirical work has often either failed to detect the operation of such benefits or shown a net cost to the presence of sexual selection. We tested whether sexual selection can increase the speed with which a conditionally deleterious allele is removed from a laboratory population of Drosophila melanogaster. The alcohol dehydrogenase null allele ( Adh –) confers slightly lower viability than wild-type alleles in the absence of ethanol but is lethal in homozygotes when ethanol comprises 6% of the medium. We tracked the frequency of this allele in artificially constructed populations reared at three different levels of ethanol (0%, 2%, and 4%) that either experienced sexual selection or did not. Loss of the deleterious Adh – allele was more rapid when sexual selection was allowed to act, especially in the presence of ethanol. We also quantified the strength of both nonsexual and sexual selection against the Adh – allele using maximum-likelihood estimation. In contrast to recent experiments employing monogamy/polygamy designs, our results demonstrate a fitness benefit to sexual selection. This is consistent with the operation of good-genes female choice.